Basal cell

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Overview

A microscopic image of a prostate gland featuring an example of basal cells Prostate gland microanatomy.png
A microscopic image of a prostate gland featuring an example of basal cells

A basal cell is a general cell type that is present in many forms of epithelial tissue throughout the body. Basal cells are located between the basement membrane and the remainder of the epithelium, effectively functioning as an anchor for the epithelial layer and an important mechanism in the maintenance of intraorgan homeostasis.

Contents

Basal cells can interact with surrounding cells including neurons, the basement membrane, columnar epithelium, and underlying mesenchymal cells. They also engage in interactions with dendritic, lymphocytic, and inflammatory cells, with the majority of these interactions occurring in the lateral intercellular gap between basal cells. [1]

Basal cells have important health implications since the most common types of skin cancer are basal cell and squamous cell carcinomas. More than 1 million instances of these cancers, referred to as non-melanoma skin cancers (NMSC) are expected to be diagnosed in the United States each year, and the incidence is rapidly increasing. Basal and squamous cell malignancies, while seldom metastatic, can cause significant local damage and disfigurement, affecting large sections of soft tissue, cartilage, and bone. [2]

Location

A microscopic image of the human trachea, showcasing the typical location of basal cells (B) between the basement membrane (BM) and the remaining epithelium Masson's trichrome staining of the Human trachea.jpg
A microscopic image of the human trachea, showcasing the typical location of basal cells (B) between the basement membrane (BM) and the remaining epithelium

Basal cells are located in various tissues throughout the body. They are located at the bottom of epithelial tissues, generally situated directly on top of the basal lamina, above the basement membrane and below the remainder of the epithelium. Examples include:

Structure

TEM image of the tracheal basal lamina in mice. The darkened areas with the arrows pointing to them display the location of hemidesmosomes connecting the layer of basal cells to the basal lamina. Ultrastructure of tracheal hemidesmosomes in mice.JPEG
TEM image of the tracheal basal lamina in mice. The darkened areas with the arrows pointing to them display the location of hemidesmosomes connecting the layer of basal cells to the basal lamina.

Regardless of their specific location, basal cells generally share a similar basic structure. They are all usually either cuboidal, polyhedral or pyramidal shaped cells with enlarged nuclei and minimal cytoplasm. [3] Basal cells are bound to each other by desmosomes, and to the basal lamina of the basement membrane by hemidesmosomes. These junctions help to create one tightly bound, continuous tissue layer that can endure mechanical stress and effectively function as a connection between the basement membrane and remaining epithelial tissue. [4]

Function

Basal cells serve two main functions in cells. They serve:

  1. To anchor and connect the epithelium to the basement membrane
  2. As the main stem cell population for the tissue they are found in, therefore responding to stimuli to maintain homeostasis within that tissue

While all basal cells, regardless of location, function similar in regards to anchoring the epithelium, the specific function and mechanisms of basal cells as stem cells varies by location. In general, basal cells can can function as either unipotent or multipotent stem cells.

Epidermal basal cells

A diagram displaying the layers of the epidermis, with basal cells comprising the stratum basale. Skinlayers.png
A diagram displaying the layers of the epidermis, with basal cells comprising the stratum basale.

In the epidermis, basal cells function as unipotent stem cells. [5] Found in the lowest layer of the epidermis, the stratum basale, basal cells continuously divide in order to replenish the squamous cells that make up the skin's surface. [6] Every time a basal cell divides, it creates two daughter cells, one is an identical basal cell, and the other is a new somatic cell that undergoes terminal differentiation. These cells gradually get pushed up through the layers of the epidermis by the constant proliferation of more new cells, gradually differentiating and flattening as they rise. This ultimately results in functional squamous cells on the outermost layer of the epidermis, the most abundant of which are called keratinocytes.

The continuous division of epidermal basal cells leads to complete epidermal turnover every 40-56 days in humans and every 8-10 days in mice. [7]

This process of proliferation and differentiation is regulated by multiple genetic and environmental factors including a calcium gradient, Vitamins A and D, epidermal growth factor (EGF), transcription factor p63, and transforming growth factor alpha (TGF-α). [8] [9] [10] [11]

Errors in the regulatory mechanisms of epidermal basal cells can cause a variety of acute and chronic ailments including psoriasis and basal cell carcinoma, which is the most common type of skin cancer, accounting for 80% of all skin cancer cases. [12] [13] Due to the structural importance of the epidermis, defects in basal cell proliferation and differentiation can also contribute to deformities such as cleft lips and Gorlin syndrome. [14] [15]

Respiratory basal cells

In the respiratory tract, basal cells function as multipotent stem cells, capable of replenishing all of the epithelial cell types including secretory, ciliated, and intermediate cells. They reside in the mucosal layer of the respiratory epithelium, and generally remain dormant. However, when a functional epithelial cell becomes damaged, a basal cell is activated to differentiate into the appropriate cell type and replace the damaged cell. [16] [17]

A diagram of cells in the respiratory epithelium. Basal cells are shown in purple, ciliated cells shown in brown, goblet cells are shown in green, and submucosal (below the epithelium) glands are shown in blue. Respiratory Tract Cells.png
A diagram of cells in the respiratory epithelium. Basal cells are shown in purple, ciliated cells shown in brown, goblet cells are shown in green, and submucosal (below the epithelium) glands are shown in blue.

In addition to functioning as stem cells, there is novel evidence to suggest that undifferentiated basal cells also contribute immune functions of the respiratory epithelium by secreting RNase. This function helps to preserve the immune capabilities of the respiratory epithelium even when it is damaged and in the process of being repaired. [18]

In the respiratory epithelium, there exists a layer of intermediate cells between the basal and differentiated cells. These intermediate cells exist in a transient state. They have begun the process of differentiation, but are not yet terminally differentiated, and as such can differentiate as needed, but have limited proliferative capacity. They play an important role in ensuring that the epithelium can be quickly repaired in response to damage. [19]

The process of respiratory basal cell differentiation is regulated by multiple factors including transcription factors such as FOXJ1, FOXA3, Sox2, and p53, proteins such as LEF-1, and interleukins IL-1α and IL-33, as well as other other cytokines. [20] [21] [22] However, the primary control of basal respiratory cell differentiation is the Notch signaling pathway, which is the main determinant of what the basal cell differentiates into. [23] High levels of NOTCH activity leads to differentiation into a secretory cell, whereas low levels lead to differentiation into a ciliated cell.

Gastrointestinal basal cells

The gastrointestinal tract consists of the esophagus, stomach, small intestines, and large intestines, and each layer is lined with distinct yet similar epithelium that necessarily contains basal cells. While the general function of these basal cells is similar throughout the entire tract, the specific mechanisms, functions, and products of these cells can vary depending on which layer the cells are located in. For example, while basal cells in both the esophagus and the stomach function as multipotent progenitor cells, they are fundamentally different because the esophageal basal cells exist as a part of a stratified squamous epithelium, whereas the gastric basal cells exist as a part of a simple columnar epithelium. Functionally, this means that since a simple epithelium is only one cell thick, differentiated cells must diffuse along the plane of the basement membrane rather than vertically through the rest of the epithelium. Furthermore, the actual products of these cells vary substantially, as esophageal basal cells mainly produce squamous epithelial cells, which function as a passive physical barrier between the lumen of the esophagus and underlying tissues, but gastric basal cells differentiate into a variety of secretory and absorptive cells that provide the main functions of the stomach including absorptive cells, chief cells, and parietal cells. [24]

Diagram and histological examples showing the structure of gastric glands in different areas of the stomach. Note how the progenitor cells (which in this case is a term synonymous with basal cells) are located in the isthmus region of the pit in both examples, and how there is a single layer of cells that comprises the epithelium. Glandulas gastricas estruct.jpg
Diagram and histological examples showing the structure of gastric glands in different areas of the stomach. Note how the progenitor cells (which in this case is a term synonymous with basal cells) are located in the isthmus region of the pit in both examples, and how there is a single layer of cells that comprises the epithelium.

In the stomach, basal cells are generally located in the isthmus region, or near the top, of gastric glands, a location that allows them to easily differentiate within the gland and then diffuse bi-directionally as they differentiate, going either to the above gastric pit or the base of the gastric gland to replenish damaged cells. Due to the harsh environment created by the acidic interior of the stomach, the basal cells propagate continuously, relying on a variety of pathways and signaling molecules to communicate what type of cells have been damaged and need to be replaced. These regulators of proliferation and differentiation include the protein Sox9, the Wnt and Notch signaling pathways, BMP's 2, 4, and 7 (which can all function as tumor suppressors), and EGF. [25] [26] [27] [28] These processes exist in a delicate state, and any errors in or disruptions of these pathways can cause a variety of ailments. For example, a Helicobacter pylori infection can cause an overexpression of EGF which leads to excessive differentiation of basal cells into gastrin cells, which in turn can lead to atrophic gastritis, a well studied precursor to gastric cancer. [29] Furthermore, if the genes coding for Jag1 or Jag2 are mutated or deleted, this can cause a disruption of the critical Notch signaling pathway, which can in turn cause uncontrolled and unregulated growth and differentiation leading to tumorigenesis. [30]

Diagram showing the location of intestinal crypts and the gradual migration of differentiating cells out of the crypt. Epithelial cell migration.tif
Diagram showing the location of intestinal crypts and the gradual migration of differentiating cells out of the crypt.

Similar to gastric basal cells, intestinal basal cells are continuously propagating. In fact, due to the vital role that the small intestine plays in nutrient absorption, basal cells in the small intestine exhibit the highest turnover rate of any cells in the body, creating an entirely new epithelium approximately every 5-7 days. [31] [32] Within the intestines, basal cells are located at the base of intestinal invaginations known as crypts, where they are nourished and protected by paneth cells and the surrounding microenvironment. These basal cells then function as multipotent progenitors, capable of differentiation into six distinct cell types, regulated by mechanisms very similar to those seen in other gastrointestinal basal cells. [33] As the cells differentiate, they migrate out from the crypt towards the lumen, until eventually dying and being released into the intestinal lumen, only to soon be replaced by a new cell.

Related Research Articles

<span class="mw-page-title-main">Skin</span> Soft outer covering organ of vertebrates

Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation.

<span class="mw-page-title-main">Integumentary system</span> Skin and other protective organs

The integumentary system is the set of organs forming the outermost layer of an animal's body. It comprises the skin and its appendages, which act as a physical barrier between the external environment and the internal environment that it serves to protect and maintain the body of the animal. Mainly it is the body's outer skin.

<span class="mw-page-title-main">Tissue (biology)</span> Group of similar cells performing a specific function

In biology, tissue is an assembly of similar cells and their extracellular matrix from the same embryonic origin that together carry out a specific function. Tissues occupy a biological organizational level between cells and a complete organ. Accordingly, organs are formed by the functional grouping together of multiple tissues.

<span class="mw-page-title-main">Epithelium</span> Tissue lining the surfaces of organs in animals

Epithelium or epithelial tissue is a thin, continuous, protective layer of cells with little extracellular matrix. An example is the epidermis, the outermost layer of the skin. Epithelial (mesothelial) tissues line the outer surfaces of many internal organs, the corresponding inner surfaces of body cavities, and the inner surfaces of blood vessels. Epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. These tissues also lack blood or lymph supply. The tissue is supplied by nerves.

<span class="mw-page-title-main">Keratinocyte</span> Primary type of cell found in the epidermis

Keratinocytes are the primary type of cell found in the epidermis, the outermost layer of the skin. In humans, they constitute 90% of epidermal skin cells. Basal cells in the basal layer of the skin are sometimes referred to as basal keratinocytes. Keratinocytes form a barrier against environmental damage by heat, UV radiation, water loss, pathogenic bacteria, fungi, parasites, and viruses. A number of structural proteins, enzymes, lipids, and antimicrobial peptides contribute to maintain the important barrier function of the skin. Keratinocytes differentiate from epidermal stem cells in the lower part of the epidermis and migrate towards the surface, finally becoming corneocytes and eventually being shed, which happens every 40 to 56 days in humans.

<span class="mw-page-title-main">Epidermis</span> Outermost of the three layers that make up the skin

The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water released from the body into the atmosphere through transepidermal water loss.

<span class="mw-page-title-main">Skin condition</span> Any medical condition that affects the integumentary system

A skin condition, also known as cutaneous condition, is any medical condition that affects the integumentary system—the organ system that encloses the body and includes skin, nails, and related muscle and glands. The major function of this system is as a barrier against the external environment.

<span class="mw-page-title-main">Hemidesmosome</span> Structures connecting keratinocyte cells to the extracellular matrix

Hemidesmosomes are very small stud-like structures found in keratinocytes of the epidermis of skin that attach to the extracellular matrix. They are similar in form to desmosomes when visualized by electron microscopy; however, desmosomes attach to adjacent cells. Hemidesmosomes are also comparable to focal adhesions, as they both attach cells to the extracellular matrix. Instead of desmogleins and desmocollins in the extracellular space, hemidesmosomes utilize integrins. Hemidesmosomes are found in epithelial cells connecting the basal epithelial cells to the lamina lucida, which is part of the basal lamina. Hemidesmosomes are also involved in signaling pathways, such as keratinocyte migration or carcinoma cell intrusion.

<span class="mw-page-title-main">Amphiregulin</span> Protein-coding gene in the species Homo sapiens

Amphiregulin, also known as AREG, is a protein synthesized as a transmembrane glycoprotein with 252 aminoacids and it is encoded by the AREG gene. in humans.

<span class="mw-page-title-main">Respiratory epithelium</span> Mucosa that serves to moisten and protect the airways

Respiratory epithelium, or airway epithelium, is a type of ciliated columnar epithelium found lining most of the respiratory tract as respiratory mucosa, where it serves to moisten and protect the airways. It is not present in the vocal cords of the larynx, or the oropharynx and laryngopharynx, where instead the epithelium is stratified squamous. It also functions as a barrier to potential pathogens and foreign particles, preventing infection and tissue injury by the secretion of mucus and the action of mucociliary clearance.

<span class="mw-page-title-main">Human skin</span> Organ covering the outside of the human body

The human skin is the outer covering of the body and is the largest organ of the integumentary system. The skin has up to seven layers of ectodermal tissue guarding muscles, bones, ligaments and internal organs. Human skin is similar to most of the other mammals' skin, and it is very similar to pig skin. Though nearly all human skin is covered with hair follicles, it can appear hairless. There are two general types of skin: hairy and glabrous skin (hairless). The adjective cutaneous literally means "of the skin".

<span class="mw-page-title-main">TP63</span> Protein-coding gene in the species Homo sapiens

Tumor protein p63, typically referred to as p63, also known as transformation-related protein 63, is a protein that in humans is encoded by the TP63 gene.

<span class="mw-page-title-main">CHUK</span> Protein-coding gene in humans

Inhibitor of nuclear factor kappa-B kinase subunit alpha (IKK-α) also known as IKK1 or conserved helix-loop-helix ubiquitous kinase (CHUK) is a protein kinase that in humans is encoded by the CHUK gene. IKK-α is part of the IκB kinase complex that plays an important role in regulating the NF-κB transcription factor. However, IKK-α has many additional cellular targets, and is thought to function independently of the NF-κB pathway to regulate epidermal differentiation.

<span class="mw-page-title-main">Epithelial cell adhesion molecule</span> Transmembrane glycoprotein

Epithelial cell adhesion molecule (EpCAM), also known as CD326 among other names, is a transmembrane glycoprotein mediating Ca2+-independent homotypic cell–cell adhesion in epithelia. EpCAM is also involved in cell signaling, migration, proliferation, and differentiation. Additionally, EpCAM has oncogenic potential via its capacity to upregulate c-myc, e-fabp, and cyclins A & E. Since EpCAM is expressed exclusively in epithelia and epithelial-derived neoplasms, EpCAM can be used as diagnostic marker for various cancers. It appears to play a role in tumorigenesis and metastasis of carcinomas, so it can also act as a potential prognostic marker and as a potential target for immunotherapeutic strategies.

miR-203

In molecular biology miR-203 is a short non-coding RNA molecule. MicroRNAs function to regulate the expression levels of other genes by several mechanisms, such as translational repression and Argonaute-catalyzed messenger RNA cleavage. miR-203 has been identified as a skin-specific microRNA, and it forms an expression gradient that defines the boundary between proliferative epidermal basal progenitors and terminally differentiating suprabasal cells. It has also been found upregulated in psoriasis and differentially expressed in some types of cancer.

Dermal fibroblasts are cells within the dermis layer of skin which are responsible for generating connective tissue and allowing the skin to recover from injury. Using organelles, dermal fibroblasts generate and maintain the connective tissue which unites separate cell layers. Furthermore, these dermal fibroblasts produce the protein molecules including laminin and fibronectin which comprise the extracellular matrix. By creating the extracellular matrix between the dermis and epidermis, fibroblasts allow the epithelial cells of the epidermis to affix the matrix, thereby allowing the epidermal cells to effectively join together to form the top layer of the skin.

Tissue engineering of oral mucosa combines cells, materials and engineering to produce a three-dimensional reconstruction of oral mucosa. It is meant to simulate the real anatomical structure and function of oral mucosa. Tissue engineered oral mucosa shows promise for clinical use, such as the replacement of soft tissue defects in the oral cavity. These defects can be divided into two major categories: the gingival recessions which are tooth-related defects, and the non tooth-related defects. Non tooth-related defects can be the result of trauma, chronic infection or defects caused by tumor resection or ablation. Common approaches for replacing damaged oral mucosa are the use of autologous grafts and cultured epithelial sheets.

Cord lining, cord tissue, or umbilical cord lining membrane, is the outermost layer of the umbilical cord. As the umbilical cord itself is an extension of the placenta, the umbilical cord lining membrane is an extension of the amniotic membrane covering the placenta. The umbilical cord lining membrane comprises two layers: the amniotic layer and the sub-amniotic layer. The umbilical cord lining membrane is a rich source of two strains of stem cells (CLSCs): epithelial stem cells (CLECs) and mesenchymal stem cells (CLMCs). Discovered by Singapore-based CellResearch Corporation in 2004, this is the best known source for harvesting human stem cells.

<span class="mw-page-title-main">Vaginal epithelium</span> Inner lining of the vagina

The vaginal epithelium is the inner lining of the vagina consisting of multiple layers of (squamous) cells. The basal membrane provides the support for the first layer of the epithelium-the basal layer. The intermediate layers lie upon the basal layer, and the superficial layer is the outermost layer of the epithelium. Anatomists have described the epithelium as consisting of as many as 40 distinct layers of cells. The mucus found on the epithelium is secreted by the cervix and uterus. The rugae of the epithelium create an involuted surface and result in a large surface area that covers 360 cm2. This large surface area allows the trans-epithelial absorption of some medications via the vaginal route.

Cédric, Baron Blanpain is a Belgian researcher in the field of stem cells. He is a tenured professor of developmental biology and genetics at Université libre de Bruxelles and director of the stem cell and cancer lab at its Faculty of Medicine. He was one of the first researchers in the world to use cell lineage tracing in cancer research and he showed for the first time the existence of cancer stem cells in solid tumors in vivo. He was selected by Nature as one of 10 People who mattered most in 2012 and he received the outstanding young investigator award of the International Society for Stem Cell Research.

References

  1. ↑ "What Are Basal and Squamous Cell Skin Cancers? | Types of Skin Cancer". www.cancer.org. Retrieved 2024-03-27.
  2. ↑ Miller, Stanley J.; Alam, Murad; Andersen, James; et al. (August 2010). "Basal Cell and Squamous Cell Skin Cancers". Journal of the National Comprehensive Cancer Network. 8 (8): 836–864. doi:10.6004/jnccn.2010.0062. ISSN   1540-1405. PMID   20870631.
  3. ↑ Haschek, Wanda M.; Witschi, Hanspeter R.; Nikula, Kristen J. (2002-01-01), Haschek, WANDA M.; Rousseaux, COLIN G.; Wallig, MATTHEW A. (eds.), "28 - Respiratory System", Handbook of Toxicologic Pathology (Second Edition), San Diego: Academic Press, pp. 3–83, doi:10.1016/b978-012330215-1/50029-6, ISBN   978-0-12-330215-1 , retrieved 2024-11-26
  4. ↑ Cote, Lauren E.; Feldman, Jessica L. (2022-06-21). "Won't You be My Neighbor: How Epithelial Cells Connect Together to Build Global Tissue Polarity". Frontiers in Cell and Developmental Biology. 10. doi: 10.3389/fcell.2022.887107 . ISSN   2296-634X. PMC   9253303 . PMID   35800889.
  5. ↑ Ghazizadeh, Soosan; Taichman, Lorne B. (2001-03-15). "Multiple classes of stem cells in cutaneous epithelium: a lineage analysis of adult mouse skin". The EMBO Journal. 20 (6): 1215–1222. doi:10.1093/emboj/20.6.1215. ISSN   0261-4189. PMC   145528 . PMID   11250888.
  6. ↑ "Basal Cell Nevus Syndrome", Encyclopedia of Cancer, Berlin, Heidelberg: Springer Berlin Heidelberg, 2011, pp. 345–346, doi:10.1007/978-3-642-16483-5_530, ISBN   978-3-642-16482-8 , retrieved 2024-03-27
  7. ↑ Koster, Maranke I. (July 2009). "Making an Epidermis". Annals of the New York Academy of Sciences. 1170 (1): 7–10. Bibcode:2009NYASA1170....7K. doi:10.1111/j.1749-6632.2009.04363.x. ISSN   0077-8923. PMC   2861991 . PMID   19686098.
  8. ↑ Fuchs, Elaine; Green, Howard (September 1981). "Regulation of terminal differentiation of cultured human keratinocytes by vitamin A". Cell. 25 (3): 617–625. doi:10.1016/0092-8674(81)90169-0. PMID   6169442.
  9. ↑ Rheinwald, James G.; Green, Howard (February 1977). "Epidermal growth factor and the multiplication of cultured human epidermal keratinocytes". Nature. 265 (5593): 421–424. Bibcode:1977Natur.265..421R. doi:10.1038/265421a0. ISSN   0028-0836. PMID   299924.
  10. ↑ Truong, Amy B.; Kretz, Markus; Ridky, Todd W.; Kimmel, Robin; Khavari, Paul A. (2006-11-15). "p63 regulates proliferation and differentiation of developmentally mature keratinocytes". Genes & Development. 20 (22): 3185–3197. doi:10.1101/gad.1463206. ISSN   0890-9369. PMC   1635152 . PMID   17114587.
  11. ↑ Barrandon, Yann; Green, Howard (September 1987). "Cell migration is essential for sustained growth of keratinocyte colonies: The roles of transforming growth factor-α and epidermal growth factor". Cell. 50 (7): 1131–1137. doi:10.1016/0092-8674(87)90179-6. PMID   3497724.
  12. ↑ Flisiak, I.; Szterling-Jaworowska, M.; Baran, A.; Rogalska-Taranta, M. (June 2014). "Effect of psoriasis activity on epidermal growth factor (EGF) and the concentration of soluble EGF receptor in serum and plaque scales". Clinical and Experimental Dermatology. 39 (4): 461–467. doi:10.1111/ced.12356. ISSN   1365-2230. PMID   24825137.
  13. ↑ "Basal Cell Carcinoma (BCC)". Yale Medicine. Retrieved 2024-11-26.
  14. ↑ Lan, Yu; Xu, Jingyue; Jiang, Rulang (2015), "Cellular and Molecular Mechanisms of Palatogenesis", Current Topics in Developmental Biology, 115, Elsevier: 59–84, doi:10.1016/bs.ctdb.2015.07.002, ISBN   978-0-12-408141-3, PMC   4663457 , PMID   26589921
  15. ↑ "Gorlin syndrome: MedlinePlus Genetics". medlineplus.gov. Retrieved 2024-11-26.
  16. ↑ Hong, Kyung U.; Reynolds, Susan D.; Watkins, Simon; Fuchs, Elaine; Stripp, Barry R. (April 2004). "In vivo differentiation potential of tracheal basal cells: evidence for multipotent and unipotent subpopulations". American Journal of Physiology-Lung Cellular and Molecular Physiology. 286 (4): L643–L649. doi:10.1152/ajplung.00155.2003. ISSN   1040-0605. PMID   12871857.
  17. ↑ Hong, Kyung U.; Reynolds, Susan D.; Watkins, Simon; Fuchs, Elaine; Stripp, Barry R. (February 2004). "Basal Cells Are a Multipotent Progenitor Capable of Renewing the Bronchial Epithelium". The American Journal of Pathology. 164 (2): 577–588. doi:10.1016/S0002-9440(10)63147-1. PMC   1602270 . PMID   14742263.
  18. ↑ Amatngalim, Gimano D.; van Wijck, Yolanda; de Mooij-Eijk, Yvonne; Verhoosel, Renate M.; Harder, Jürgen; Lekkerkerker, Annemarie N.; Janssen, Richard A. J.; Hiemstra, Pieter S. (2015-04-01). "Basal Cells Contribute to Innate Immunity of the Airway Epithelium through Production of the Antimicrobial Protein RNase 7". The Journal of Immunology. 194 (7): 3340–3350. doi:10.4049/jimmunol.1402169. ISSN   0022-1767. PMID   25712218.
  19. ↑ Boers, James E.; Ambergen, Anton W.; Thunnissen, Frederik B. J. M. (1998-06-01). "Number and Proliferation of Basal and Parabasal Cells in Normal Human Airway Epithelium". American Journal of Respiratory and Critical Care Medicine. 157 (6): 2000–2006. doi:10.1164/ajrccm.157.6.9707011. ISSN   1073-449X. PMID   9620938.
  20. ↑ "Foxj1 forkhead box J1 [Mus musculus (house mouse)] - Gene - NCBI". www.ncbi.nlm.nih.gov. Retrieved 2024-11-28.
  21. ↑ "FOXA3 forkhead box A3 [Homo sapiens (human)] - Gene - NCBI". www.ncbi.nlm.nih.gov. Retrieved 2024-11-28.
  22. ↑ Whitsett, Jeffrey A. (November 2018). "Airway Epithelial Differentiation and Mucociliary Clearance". Annals of the American Thoracic Society. 15 (Supplement_3): S143–S148. doi:10.1513/AnnalsATS.201802-128AW. ISSN   2329-6933. PMC   6322033 . PMID   30431340.
  23. ↑ Artavanis-Tsakonas, Spyros; Rand, Matthew D.; Lake, Robert J. (1999-04-30). "Notch Signaling: Cell Fate Control and Signal Integration in Development". Science. 284 (5415): 770–776. Bibcode:1999Sci...284..770A. doi:10.1126/science.284.5415.770. ISSN   0036-8075. PMID   10221902.
  24. ↑ Hayakawa, Yoku; Nakagawa, Hiroshi; Rustgi, Anil K.; Que, Jianwen; Wang, Timothy C. (2021-08-05). "Stem cells and origins of cancer in the upper gastrointestinal tract". Cell Stem Cell. 28 (8): 1343–1361. doi:10.1016/j.stem.2021.05.012. ISSN   1934-5909. PMC   8844710 . PMID   34129814.
  25. ↑ Willet, Spencer G.; Thanintorn, Nattapon; McNeill, Helen; Huh, Sung-Ho; Ornitz, David M.; Huh, Won Jae; Hoft, Stella G.; DiPaolo, Richard J.; Mills, Jason C. (2023). "SOX9 Governs Gastric Mucous Neck Cell Identity and Is Required for Injury-Induced Metaplasia". Cellular and Molecular Gastroenterology and Hepatology. 16 (3): 325–339. doi:10.1016/j.jcmgh.2023.05.009. PMC   10444955 . PMID   37270061.
  26. ↑ Kapalczynska, Marta; Lin, Manqiang; Maertzdorf, Jeroen; Heuberger, Julian; Muellerke, Stefanie; Zuo, Xiangsheng; Vidal, Ramon; Shureiqi, Imad; Fischer, Anne-Sophie; Sauer, Sascha; Berger, Hilmar; Kidess, Evelyn; Mollenkopf, Hans-Joachim; Tacke, Frank; Meyer, Thomas F. (2022-03-24). "BMP feed-forward loop promotes terminal differentiation in gastric glands and is interrupted by H. pylori-driven inflammation". Nature Communications. 13 (1): 1577. Bibcode:2022NatCo..13.1577K. doi:10.1038/s41467-022-29176-w. ISSN   2041-1723. PMC   8948225 . PMID   35332152.
  27. ↑ Wölffling, Sarah; Daddi, Alice Anna; Imai-Matsushima, Aki; Fritsche, Kristin; Goosmann, Christian; Traulsen, Jan; Lisle, Richard; Schmid, Monika; Reines-Benassar, Maria del Mar; Pfannkuch, Lennart; Brinkmann, Volker; Bornschein, Jan; Malfertheiner, Peter; Ordemann, Jürgen; Link, Alexander (August 2021). "EGF and BMPs Govern Differentiation and Patterning in Human Gastric Glands". Gastroenterology. 161 (2): 623–636.e16. doi:10.1053/j.gastro.2021.04.062. ISSN   0016-5085. PMID   33957136.
  28. ↑ Shirai, Yo-taro; Ehata, Shogo; Yashiro, Masakazu; Yanagihara, Kazuyoshi; Hirakawa, Kosei; Miyazono, Kohei (December 2011). "Bone Morphogenetic Protein-2 and -4 Play Tumor Suppressive Roles in Human Diffuse-Type Gastric Carcinoma". The American Journal of Pathology. 179 (6): 2920–2930. doi:10.1016/j.ajpath.2011.08.022. PMC   3260801 . PMID   21996676.
  29. ↑ Uemura, Naomi; Okamoto, Shiro; Yamamoto, Soichiro; Matsumura, Nobutoshi; Yamaguchi, Shuji; Yamakido, Michio; Taniyama, Kiyomi; Sasaki, Naomi; Schlemper, Ronald J. (2001-09-13). "Helicobacter pylori Infection and the Development of Gastric Cancer". New England Journal of Medicine. 345 (11): 784–789. doi:10.1056/NEJMoa001999. ISSN   0028-4793. PMID   11556297.
  30. ↑ Huang, Haidi; Jiang, Yu; Liu, Jiangying; Luo, Dan; Yuan, Jianghong; Mu, Rongzi; Yu, Xiang; Sun, Donglei; Lin, Jihong; Chen, Qiyue; Li, Xinjing; Jiang, Ming; Xu, Jianming; Chu, Bo; Yin, Chengqian (2024-05-15). "Jag1/2 maintain esophageal homeostasis and suppress foregut tumorigenesis by restricting the basal progenitor cell pool". Nature Communications. 15 (1): 4124. Bibcode:2024NatCo..15.4124H. doi:10.1038/s41467-024-48347-5. ISSN   2041-1723. PMC   11096375 . PMID   38750026.
  31. ↑ van der Flier, Laurens G.; Clevers, Hans (2009-03-01). "Stem Cells, Self-Renewal, and Differentiation in the Intestinal Epithelium". Annual Review of Physiology. 71 (1): 241–260. doi:10.1146/annurev.physiol.010908.163145. ISSN   0066-4278. PMID   18808327.
  32. ↑ "Understanding how the intestine replaces and repairs itself". hsci.harvard.edu. Retrieved 2024-12-06.
  33. ↑ Gehart, Helmuth; Clevers, Hans (January 2019). "Tales from the crypt: new insights into intestinal stem cells". Nature Reviews Gastroenterology & Hepatology. 16 (1): 19–34. doi:10.1038/s41575-018-0081-y. ISSN   1759-5053. PMID   30429586.
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